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BIOL2007 Evolution of Warning Colour and Mimicry DEFENSIVE BIOL2007 Painted palatable male day flying moths Evolution of warning colour and mimicry (Callosamia promethea), a natural mimic of the unpalatable papilionid Battus philenor, to resemble DEFENSIVE COLORATION unpalatable and palatable species [OVERHEAD 6A]. Camouflage - also called crypsis, cryptic coloration. Recaptured by baiting traps with females, which "call" using pheromones. Flash coloration. e.g. flying grasshoppers which have blue or red underwings or Mimics of palatable species recaptured at a rate “underwing” moths. Eyed hawkmoth. only 40% of rate of recapture of mimics of unpalatable species Aposematism or warning colour. Avertises ˆ s=0.6. punishment could be in the form of: Similar field experiments with Müllerian mimicry in Heliconius: Unpalatability e.g. wasp stings, distastefulness. s .0.2-0.6. Smelliness e.g. skunk. Selection for mimicry can be strong! Dangerousness e.g. coral snakes (cobra family). Evolution of unpalatability and warning colour Warning colours are usually learned Unpalatability But innate recognition of coral snake patterns. Potentially a costly altruism (a) metabolic energy to sequester or synthesize the Mimicry. A mimic evolves to look like unpalatable compounds an unpleasant model species. (b) danger in teaching predators Two major kinds of mimicry are: Benefit to the individual potentially enormous Batesian mimicry - (Bates 1862), e.g compounds are sequestered wings of Danaidae cheats, parasites! Müllerian mimicry - after Müller If the cost > the individual benefit (1879), mutualists. unpalatability would be a true altruism EVIDENCE FOR MIMICRY AND a warningly coloured cheat would do rather well by WARNING COLOUR avoiding the cost of chemical sequestration, but gaining protection due to others' unpalatability. Experimental evidence for the efficacy of warning colour and mimicry. Kin selection a possible mode of evolution. Jane van Zandt Brower & Lincoln Brower in the Fisher (1930) suggested that this is why gregarious 1950s performed laboratory experiments with larvae, usually laid as eggs by a single female, blue jays and monarch butterflies (Danaus associated with warning colour plexippus): If Benefits to the individual > costs, unpalatability 1) Jays fed monarchs became sick & not altruism can evolve under individual selection wiped beak alone. (possibly helped along by kin selection) ˆ monarchs are unpalatable. 2) Jays learn to avoid monarchs Solitary larvae, like the famous monarch butterfly ˆ appropriate learning is possible Danaus plexippus 3) Having learnt, Jays avoid mimics of monarchs Adults migrate thousands of km as though as far ˆ mimicry theory correct away from their relatives as possible! EVIDENCE FOR MIMICRY AND Living gregariously. WARNING COLOUR Experimental evidence for the efficacy of So why do some unpalatable caterpillars live in warning colour and mimicry. family groups? Field experiments few! e.g. Jeffords, Surely this is good evidence for kin selection! No! Waldbauer & Sternberg - Batesian mimicry. 1 Some very good selfish reasons why unpalatable EVOLUTION OF MIMICRY species should live groups: Punnett early 20th C., a Mendelian – Darwinian evolution did not explain mimicry Predator satiation by groups. Group defence, coordinated signalling. Fisher 1930 proposed gradualistic explanation By aggregating, avoid some predators entirely. Goldschmidt in 1940-1945, proposed "systemic mutation" - single, massive chromosomal mutations EVOLUTION OF WARNING COLOURS that reorganized the whole genome. Suggested that after evolution of unpalatability homologous developmental pathways were used in model and mimic. Modern view: a little of both! Batesian mimicry Sir Cyril Clarke and Philip Sheppard: Papilionidae. Goldschmidt was wrong on the details.e.g. Papilio memnon 1) Mimics: analogous resemblance 2) Mimicry not inherited as a single gene: Major locus + unlinked components Major locus itself is “supergene” 3) Polymorphism in Batesian mimicry if tight linkage disequilibrium at “supergene” shifting balance/kin selection Müllerian mimicry individual selection preadaptation to signalling: Selection AGAINST rarity. Few polymorphisms sexual selection Therefore, do not expect "supergenes" flash coloration In Heliconius a number of unlinked loci Batesian mimicry (warning On the other hand, mimicry genes do have large colour before unpal.) effects, with strong selection † Fisher gradualist Müllerian mimicry theory. sensory bias e.g. Dry gene in Heliconius erato Nicholson “two-step” model of Müllerian mimicry: Evidence 1) Mimicry provides a "rugged" adaptive landscape diversity of warning patterns within unpalatable 2) A major mutation does help Müllerian adaptation clades: suggests warning colour evolves rapidly. 3) More minor genes do “Tidying up” MIMICRY AND WARNING COLOUR Provides examples of more general evolutionary thinking: Social evolution and kin selection Linkage disequilibria and evolution at >1 gene Evolutionary developmental genetics Drift/shifting balance in evolution Race formation and speciation Biogeography 2.
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